Recovery and Decline of Lake Whitefish in U.S. Waters of Eastern Lake Ontario,

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1 University of Nebraska - Lincoln DigitalCommons@University of Nebraska - Lincoln Publications, Agencies and Staff of the U.S. Department of Commerce U.S. Department of Commerce 2005 Recovery and Decline of Lake Whitefish in U.S. Waters of Eastern Lake Ontario, Randall W. Owens U.S. Geological Survey Robert O'Gorman U.S. Geological Survey, rogorman@usgs.gov Thomas H. Eckert Cape Vincent Fisheries Research Station Brian F. Lantry U.S. Geological Survey Dawn E. Dittman U.S. Geological Survey Follow this and additional works at: Part of the Environmental Sciences Commons Owens, Randall W.; O'Gorman, Robert; Eckert, Thomas H.; Lantry, Brian F.; and Dittman, Dawn E., "Recovery and Decline of Lake Whitefish in U.S. Waters of Eastern Lake Ontario, " (2005). Publications, Agencies and Staff of the U.S. Department of Commerce This Article is brought to you for free and open access by the U.S. Department of Commerce at DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in Publications, Agencies and Staff of the U.S. Department of Commerce by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln.

2 Recovery and Decline of Lake Whitefish in U.S. Waters of Eastern Lake Ontario, Randall W. Owens U.S. Geological Survey Great Lakes Science Center Lake Ontario Biological Station 17 Lake Street Oswego, New York, U.S.A Robert O Gorman 1 U.S. Geological Survey Great Lakes Science Center Lake Ontario Biological Station 17 Lake Street Oswego, New York, U.S.A Thomas H. Eckert New York State Department of Environmental Conservation Cape Vincent Fisheries Research Station Cape Vincent, New York, U.S.A Brian F. Lantry U.S. Geological Survey Great Lakes Science Center Lake Ontario Biological Station 17 Lake Street Oswego, New York, U.S.A Dawn E. Dittman U.S. Geological Survey U.S. Geological Survey Great Lakes Science Center Tunison Laboratory of Aquatic Sciences 3075 Gracie Rd. Cortland, New York, U.S.A Corresponding author: rogorman@usgs.gov 141 Mohr, L.C., and Nalepa, T.F. (Editors) Proceedings of a workshop on the dynamics of lake whitefish (Coregonus clupeaformis) and the amphipod Diporeia spp. in the Great Lakes. Great Lakes Fish. Comm. Tech. Rep. 66.

3 Abstract The lake whitefish (Coregonus clupeaformis) was an important member of the native fish community and a valued commercial species in Lake Ontario. Lake whitefish were common in U.S. waters of the lake until 1965 and very abundant in Canadian waters through the early 1970s, although their numbers declined shortly thereafter. During , lake whitefish stocks remained depressed throughout the lake as a result of the combined effects of degraded water quality, overfishing, and predation. Rainbow smelt (Osmerus mordax) probably preyed on whitefish fry, and sea lamprey (Petromyzon marinus) preyed on adults. During , lake whitefish stocks began to recover in eastern Lake Ontario, and their buildup continued into the mid-1990s. Reasons for the recovery likely included control of the sea lamprey population and a reduction in the number of piscivorous rainbow smelt. By 1997, lake whitefish abundance had declined severely again; some fish appeared to have dispersed from the northeastern to the southeastern regions of the lake, and the depth of capture increased. We believe that the collapse of Diporeia spp. populations during was responsible for the decline in the lake whitefish populations and the shifts in geographic and bathymetric distribution because lake whitefish fed primarily on Diporeia spp. After the collapse of Diporeia spp. populations, lake whitefish in southeastern Lake Ontario fed on Mysis relicta and quagga mussels (Dreissena bugensis). Changing from a diet of high-lipid Diporeia spp. to low-lipid dreissenids and foraging on Mysis relicta at lower temperatures are apparently hampering the rebuilding of lake whitefish stocks. Introduction Historically, lake whitefish (Coregonus clupeaformis, hereafter, whitefish) were an abundant and important commercial species throughout Lake Ontario (Rathbun and Wakeham 1897; Koelz 1926). Commercial landings 142

4 of whitefish in U.S. waters between 1885 and 1965 ranged from 9-91 mt per year and were roughly 10 times that amount in Canadian waters. By 1966, annual whitefish landings had dropped to <1 mt from U.S. waters. By 1975, the U.S. commercial fishery no longer existed, and whitefish landings had plummeted to 4.5 mt in Canadian waters (Baldwin et al. 1979). The reasons for the decline of whitefish stocks in Lake Ontario likely included predation by sea lamprey (Petromyzon marinus) (Christie 1973), overfishing, predation on young-of-the-year (YOY) whitefish by the exotic rainbow smelt (Osmerus mordax, hereafter, smelt) (Warner and Fenderson 1963; Bergstedt 1983; Evans and Loftus 1987), and perhaps degradation of water quality (Christie 1973; Hoyle et al., 2003; Owens et al., 2003). During , whitefish were conspicuously absent in bottom trawl surveys conducted jointly by the U.S. Geological Survey (USGS) and the New York State Department of Environmental Conservation (NYSDEC) in U.S. waters of Lake Ontario (Owens et al., 2003). Whitefish stocks began to recover in eastern Lake Ontario during , and recovery was well under way by 1995 (Hoyle et al., 2003; Owens et al., 2003). In June of 1997, however, off Prince Edward County, Ontario, Canada, emaciated and dead whitefish were observed (suggestive of starvation), and whitefish recruitment declined, which eventually led to a decline in abundance (Hoyle et al. in press). In 1996, whitefish abundance declined near Henderson Harbor, New York, located about 50-km southeast of Prince Edward County. In 1997, an unusually large number of whitefish (about 70) were caught approximately 50 km southwest of Henderson Harbor near Oswego, New York, suggesting that whitefish might have been dispersing from the shallow northeastern basin of the lake. These recent changes in abundance, bathymetric distribution, and lengthweight relationships of whitefish were likely the result of the collapse of the burrowing amphipod Diporeia spp., (hereafter diporeia as a common name) population, which began as early as 1992 in eastern Lake Ontario (Owens and Dittman 2003) and spread lakewide thereafter (Dermott 2001; Lozano et al. 2001). Diporeia was the staple item in the diet of whitefish (Hart 1931; Hoyle et al., 2003; Owens and Dittman 2003). Its population collapse coincided with the establishment and expansion of the exotic zebra mussel (Dreissena polymorpha) and quagga mussel (D. bugensis), first noted in Lake Ontario in 1989 (Mills et al. 1999). In this paper, we describe the rise and fall in whitefish abundance, dispersion from the Henderson Harbor area, 143

5 changes in bathymetric distribution, and changes in their length-weight regressions in U.S. waters of eastern Lake Ontario during We also discuss the events that were likely responsible for the changes. Methods Determination of whitefish abundance in followed procedures outlined in Owens et al. (2003). Briefly, abundance was the total number caught each year during four separate bottom-trawl surveys adjusted to a standard effort of 350 tows, which is very close to the average number of tows conducted annually since Trawl surveys were conducted from mid-april to early November in U.S. waters. The number of locations fished in each survey varied from 6-13, depending on the targeted species. Trawl locations were spaced at about 25-km intervals along the southern and eastern shores of Lake Ontario, starting near Olcott and finishing near the head of the St. Lawrence River (Fig. 1). Because the abundance of whitefish varied geographically, we divided the lake into two regions: eastern and western. The eastern region was further divided into two areas: Henderson Harbor (1, Fig. 1), an area encompassing historically important whitefish grounds where whitefish were most abundant during this study, and Oswego (2, Fig. 1), a transition area where whitefish abundance spiked temporarily in The western and largest region (3, Fig. 1), where whitefish abundance was uniformly low during , extended from Sodus westward to the Niagara River. Four to eight locations were fished within the western region, depending on the targeted species. About nine tows were made at each location covering depths from m with the exception of the Henderson Harbor area where the maximum depth is about 55 m. 144

6 Fig. 1. Map of Lake Ontario depicting three geographic areas of varying whitefish abundance. The eastern region comprises two areas: Henderson Harbor (1) and Oswego (2). The western region (3) extends from Sodus westward to the Niagara River. The trawls used by the USGS and NYSDEC are constructed identically, as are the trawl doors and harness riggings. After 1995, with the exception of the autumn survey that targeted slimy sculpins (Cottus cognatus), the USGS 12-m bottom trawl was replaced with a three-seam bottom trawl featuring small rollers to counteract heavy fouling by the rapidly increasing population of dreissenids. Trawling depths remained the same as in previous surveys. The new trawl was calibrated against the old trawl by making over 100 side-by-side tows. In theory, whitefish should be more vulnerable to the three-seam bottom trawl because the three-seam bottom trawl s opening is twice as high as that of the 12-m trawl, although its spread is similar. The 145

7 three-seam trawl was towed at the same speed as the old trawl. During our calibration trials, 17 whitefish were caught in the three-seam trawl compared with 10 whitefish in the 12-m bottom trawl. We did not attempt to calculate a conversion factor because no whitefish were taken in either net in over 95% of the paired tows. Therefore, we assumed that whitefish were similarly vulnerable to the old and new gear. All whitefish were measured (total length, mm) and weighed (nearest g) shortly after capture. We used analysis of covariance (ANCOVA) to test for differences in lengthweight regressions for mature whitefish ( 300 mm) during two time periods: pre-dreissenid colonization ( ) and post-dreissenid colonization ( ). We used a log 10 transformation to linearize the data. If the ANCOVA indicated that the slopes were equal, we used least-squares means to test for differences in elevation of regression lines. The first dreissenids were caught in our trawls in 1991, which is the year we used to separate the two time periods (pre- and post-colonization). The recovery of whitefish may be related to the reduction of large smelt through predation by salmonines. To determine if the resurgence of whitefish followed or preceded the decline in abundance of smelt, we calculated the abundance of large ( 150-mm) smelt from the average number caught per 10-min tow during our joint bottom-trawl surveys conducted annually in June, To obtain the total catch of large smelt, the number of all smelt caught in one tow was multiplied by the proportion of smelt 150 mm from length-frequencies for smelt from nearly all trawl tows. We correlated the total number of all six species of salmonids stocked two years before (after two years in the lake, these fish should have been of sufficient size to eat all sizes of smelt) with the abundance of large smelt in U.S. waters during For example, the number of trout and salmon stocked in 1978 (1.53 million) was paired with the number of large smelt caught per trawl tow in 1980 (141). We limited the correlation of smelt and salmonines to to eliminate any effects on smelt due to dreissenid colonization (when dreissenid became abundant after 1992, water clarity increased and the bathymetric distribution of smelt shifted, potentially increasing the vulnerability of smelt to predation). 146

8 Results Whitefish were uncommon near Henderson Harbor during , but their abundance increased during (Fig. 2). About 97% of all whitefish caught in U.S. waters came from Henderson Harbor even though effort in this area was only about 15% of the total. This increase in whitefish numbers followed a decline in abundance of large smelt (Fig. 2). The decline in the abundance of large smelt during was significantly correlated (r = 0.802, P = 0.001) with the number of trout and salmon stocked two years prior in U.S. waters (Fig. 2). Abundance of whitefish began to decline sharply near Henderson Harbor beginning in Between 1995 and 2001, the number of whitefish caught in this area dropped by more than 95%. Seventy whitefish were caught near Oswego in the autumn of 1997 whereas whitefish were rarely caught there during Abundance of whitefish near Oswego declined sharply after 1997 (Fig. 2). Catches of whitefish in the western region were low during even though 65% of all trawling effort occurred in this large area. Contrary to the trend toward increasing abundance in the Henderson Harbor region during , the trend in southwestern Lake Ontario was low catches, averaging <1% of the total. 147

9 Fig. 2. Total number of whitefish caught per 350 bottom-trawl tows. HEN = Henderson Harbor area (black bars), OSW = Oswego area (cross-hatched bars), Big Smelt = mean number of large ( 150 mm) rainbow smelt (white bars) per tow in U.S. waters of Lake Ontario, and Salmonids = the number stocked (other than fry) two-years before in U.S. waters of Lake Ontario. Our ANCOVA indicated that the slopes for the length-weight regressions for whitefish were not different (P = >0.05). The least-squares means were significantly different, indicating that the regression line from postdreissenid colonization was below that of the pre-dreissenid time period (P = 0.006) (Fig. 3). For example, the predicted weight of a 500-mm whitefish declined by nearly 20%: from 1216 g during (log 10 W = log 10 L) to 973 g during (log 10 W = log 10 L). 148

10 Fig. 3. Length-weight regressions for whitefish in Lake Ontario prior to dreissenid establishment ( , open circles with regression line) and after dreissenid establishment ( , solid triangles with regression line). Discussion Although whitefish were probably common throughout Lake Ontario prior to 1900, they were likely most abundant in the northeastern region of the lake (Rathbun and Wakeham 1897; Koelz 1926). This region, especially from Henderson Harbor northwards toward Prince Edward County, is shallow (<60 m) with an extensive area between the 20- and 50-m depth contours, has suitable temperature regimes for whitefish, and the region has numerous islands that protect whitefish spawning shoals from intense wave action. Whitefish were much more abundant in Canadian than in U.S. waters, where landings were one-tenth lower (Baldwin et al. 1979). Landings were essentially nil after 1965 in U.S. waters and after 1974 in Canadian waters, suggesting that the lakewide population had nearly 149

11 vanished. Our assessment fishing showed that whitefish stocks in U.S. waters remained depressed during The stocks near Henderson Harbor began to recover around 1987 shortly after recovery began off Prince Edward County (Hoyle et al. 2003), and whitefish abundance in these areas was high by Possible reasons for this recovery include sea lamprey control, which began in the late 1970s (Schneider et al. 1996), and a reduction in the abundance of large smelt because of increased predation by the expanding lake trout (Salvelinus namaycush) and salmon (Oncorhynchus spp.) populations. Hatchery-reared trout and salmon consume smelt (Brandt 1986; Elrod and O Gorman 1991; J. R. Lantry 2001). Some studies suggest a negative relation between smelt abundance and whitefish reproductive success a result of smelt preying on YOY whitefish (Warner and Fenderson 1963; Evans and Loftus 1987). Yearling smelt are able to consume fish larvae but piscivory on advanced fry or small juveniles (40-75 mm) is generally greatest among smelt >150 mm (O Gorman 1974; Evans and Loftus 1987). Casselman et al. (1996) attributed the population recovery of whitefish near Prince Edward County to a decline in the numbers of large smelt and to temperatures more favorable for whitefish egg incubation. By 1997, however, whitefish stocks in U.S. waters were showing signs of stress. Whitefish were scarcer near Henderson Harbor where they had been historically abundant, and part of the population may have dispersed southward along the eastern shore, as suggested by a sharp increase in abundance near Oswego in After 1997, whitefish were again uncommon in U.S. waters. Length-weight regressions for whitefish indicated that, for a given length, they were 20% lighter after Hoyle et al. (2003) also noted a decline in growth of whitefish off Prince Edward County after Diporeia was formerly the principal prey in whitefish diet (Hart 1931) and the most abundant benthic macroinvertebrate in the open waters of the lake (Golini 1979). Diporeia is much higher in lipid content than many other macroinvertebrates, including dreissenids (Gardner et al. 1985; Nalepa et al. 1993). The decline in diporeia was first detected near Oswego in southeastern Lake Ontario in autumn 1992 at depths beyond 60 m (Owens and Dittman 2003). By 1995, diporeia had disappeared, or nearly disappeared, from a wide area off of Prince Edward County and the Bay of Quinte (Dermott 2001), the same area where Hoyle et al. (2003) observed dead and emaciated whitefish in By 1999, the diporeia population had 150

12 collapsed along the southern and eastern shores of the lake at depths <100 m and in some areas at depths >100 m (Dermott 2001; Lozano et al. 2001; Owens and Dittman 2003). In 1992, near Olcott, diporeia densities were as high as 8000 m - ² at depths of 55 and 75 m; by 1999, densities had declined to or near zero (Owens and Dittman 2003). Whitefish collected from deep (>70 m) U.S. waters consumed mostly Mysis relicta, sphaeriids (fingernail clams), and quagga mussels (Owens and Dittman 2003). During 1998, about 75% of the whitefish collected for diet analysis had Mysis relicta in the gut, about 95% had fingernail clams, and about 15% had quagga mussels. Only a few whitefish consumed diporeia in 1998, and none were found during During , more than 90% of the whitefish consumed Mysis relicta, about 95% consumed fingernail clams, and about 40% consumed quagga mussels. In U.S. waters, the mean depth of capture increased from about 30 m to nearly 80 m, reflecting perhaps a search for alternative prey (Owens et al. 2003). Whitefish are now foraging where temperatures are much colder (Owens et al. 2003). This change in foraging strategy was apparently not sufficient to stave off a further decline in abundance in eastern Lake Ontario. The demise of the whitefish populations in Lake Ontario may be indirectly linked to colonization by dreissenids. The diporeia population collapsed following the rapid expansion of the dreissenid population during (Dermott 2001; Lozano et al. 2001). Although no cause-and-effect relationship has been identified between dreissenids and diporeia, one hypothesis points to competition between dreissenids and diporeia for algae, especially diatoms (Dermott and Kerec 1997; Nalepa et al. 1998). Diatoms and bacteria have been identified as important components of the diet of diporeia (Marzolf 1965). The collapse of the diporeia population could be unrelated to dreissenid establishment and perhaps is due to an exotic pathogen, climate change, or other factors. We do not expect the whitefish populations to recover unless the diporeia population either recovers or is replaced by another suitable prey. Also, the establishment of another exotic species, the round goby (Neogobius melanostomus), may further impede a whitefish recovery. Gobies were found to suppress the recruitment of the mottled sculpin (Cottus bairdi) in 151

13 Lake Michigan (Janssen and Jude 2001) and to prey on small fishes in the St. Clair River, Michigan (French and Jude 2001). Acknowledgements We thank two anonymous reviewers for their constructive comments. This article is Contribution 1227 of the USGS Great Lakes Science Center. References Baldwin, N.S., Saalfeld, R.W., Ross, M.A., and Buettner, H.J Commercial fish production in the Great Lakes. Great Lakes Fish. Comm. Tech. Rep. No. 3. Bergstedt, R.A Origins of rainbow smelt in Lake Ontario. J. Great Lakes Res. 4: Brandt, S.B Food of trout and salmon in Lake Ontario. J. Great Lakes Res. 12: Casselman, J M., Hoyle, J.A., and Brown, D.M Resurgence of lake whitefish, Coregonus clupeaformis, in Lake Ontario in the 1980s. Great Lakes Res. Rev. 2: Christie, W.J A review of the changes in the fish species composition of Lake Ontario. Great Lakes Fish. Comm. Tech. Rep. No. 23. Dermott, R Sudden disappearance of the amphipod Diporeia from eastern Lake Ontario, J. Great Lakes Res. 27: Dermott, R., and Kerec, D Changes to the deep-water benthos of eastern Lake Erie since the invasion of Dreissena: Can. J. Fish. Aquat. Sci. 54: Elrod, J.H., and O Gorman, R Diet of juvenile lake trout in southern Lake Ontario in relation to abundance and size of prey fishes, Trans. Am. Fish. Soc. 13:

14 Evans, D.O., and Loftus, D.H Colonization of inland lakes in the Great Lakes region by rainbow smelt, Osmerus mordax: their freshwater niche and effects on indigenous fishes. Can. J. Fish. Aquat. Sci. 44 (Suppl. 2): French, J.R.P., III, and Jude, D.J Diets and diet overlap of nonindigenous gobies and small benthic native fishes co-inhabiting the St. Clair River, Michigan. J. Great Lakes Res. 27: Gardner, W.S., Nalepa, T.F., Frez, W.A., Cichocki, E.A., and Landrum, P.F Seasonal patterns in lipid content of Lake Michigan macroinvertebrates. Can. J. Fish. Aquat. Sci. 42: Golini, V.I Benthic macro- and meioinvertebrates of Lake Ontario, 1977: distribution and relative abundance. Fish. Mar. Serv. Ms. Rep. No Canada Centre for Inland Waters, Burlington, Ontario, Canada. Hart, J.L The food of the whitefish Coregonus clupeaformis (Mitchill) in Ontario waters, with a note on the parasites. Contrib. Can. Biol. Fish. No : Hoyle, J.A., Casselman, J.M., Dermott, R., and Schaner, T Resurgence and decline of lake whitefish (Coregonus clupeaformis) stocks in eastern Lake Ontario, In State of Lake Ontario: Past, present, and future. Edited by M. Munawar. Ecovision World Monograph Series. Aquatic Ecosystem Health and Management Society, Burlington, Ontario, Canada. Janssen, J., and Jude, D.J Recruitment failure of mottled sculpin Cottus bairdi in Calumet Harbor, southern Lake Michigan, induced by the newly introduced round goby Neogobius melanostomus. J. Great Lakes Res. 27: Koelz, W Fishing industry of the Great Lakes. Rep. U.S. Comm. Fish. (1925): Lantry, J.R Spatial and temporal dynamics of predation by Lake Ontario trout and salmon. Master's thesis. State University of New York, Environmental Science and Forestry. Syracuse, NY. 153

15 Lozano, S.J., Scharold, J.V., and Nalepa, T.F Recent declines in macroinvertebrate densities in Lake Ontario. Can. J. Fish. Aquat. Sci. 58: Marzolf, R.G Substrate relations of the burrowing amphipod Pontoporeia affinis in Lake Michigan. Ecology 46: Mills, E.L., Chrisman, J.R., Baldwin, B.S., Howell, T., Owens, R.W., O Gorman, R., Roseman, E.F., and Raths, M.K Changes in the dreissenid community and a shift toward dominance of the quagga mussel (Dreissena bugensis) in the lower Great Lakes. J. Great Lakes Res. 25: Nalepa, T.F., Cavaletto, J.F., Ford, M., Gordon, W.M., and Wimmer, M Seasonal and annual variation in weight and biochemical content of the zebra mussel, Dreissena polymorpha, in Lake St. Clair. J. Great Lakes Res. 19: Nalepa, T.F., Hartson, D.J., Fanslow, D.L., Lang, G.A., and Lozano, S.J Decline in benthic macroinvertebrate populations in southern Lake Michigan. Can. J. Fish. Aquat. Sci. 55: O Gorman, R Predation by rainbow smelt (Osmerus mordax) on young-ofthe-year alewives (Alosa pseudoharengus) in the Great Lakes. Prog. Fish- Cult. 36: Owens, R.W., O Gorman, R., Eckert, T.H., and Lantry, B.F The offshore fish community in southern Lake Ontario. In State of Lake Ontario: Past, present, and future. Edited by M. Munawar. Ecovision World Monograph Series. Aquatic Ecosystem Health and Management Society, Burlington, Ontario, Canada. Owens, R.W., and Dittman, D.E Shifts in the diets of slimy sculpin (Cottus cognatus) and lake whitefish (Coregonus clupeaformis) in Lake Ontario following the collapse of the burrowing amphipod, Diporeia. Aquat. Ecosyst. Health and Manage. 6: Rathbun, R., and Wakeham, W Report of the joint commission relative to the preservation of the fisheries in waters contiguous to Canada and the United States. House of Representatives, 2 nd session, Doc. No

16 Schneider, C.P., Owens, R.W., Bergstedt, R.A., and O Gorman, R Predation by sea lamprey (Petromyzon marinus) on lake trout (Salvelinus namaycush) in southern Lake Ontario, Can. J. Fish. Aquat. Sci. 53: Warner, K., and Fenderson, O.C The salmon and trout fishery of Fish River lakes, Maine. Trans. Am. Fish. Soc. 92:

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